RESEARCH ARTICLES D. King, J. Roughgarden, Theor. Popul. Biol. 21, 194 (1982). D. Cohen, J. Theor. Biol. 12, 119 (1966). D. King, J. Roughgarden, Theor. Popul. Biol. 22, 1 (1982). P. Haccou, Y. Iwasa, Theor. Popul. Biol. 47, 212 (1995). E. Kussell, S. Leibler, Science 309, 2075 (2005). L. Watson, M. J. Dallwitz, The Families of Flowering Plants: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 29 July 2006 http://delta-intkey.com (1992 onward). 34. T. Keller, J. Abbott, T. Moritz, P. Doerner, Plant Cell 18, 598 (2006). 35. A. C. Whibley et al., Science 313, 963 (2006). 36. S. Gavrilets, Fitness Landscapes and the Origin of Species, S. Levin, H. Horn, Eds., Monographs in Population Biology (Princeton Univ. Press, Princeton and Oxford, 2004). 37. We thank J. Avondo for help with visualizing 3D fitness landscapes, M. Clauss for helpful discussions on bet-hedging, and C. Thébaud for advice on inflorescence databases. This research was funded by grants from Human Frontier Science Program (E.C. and P.P.), Natural Sciences and Engineering Research Council of Canada (P.P. and L.H.), and the Biotechnology and Biological Sciences Research Council, UK (E.C.). Marine Radiocarbon Evidence for the Mechanism of Deglacial Atmospheric CO2 Rise Thomas M. Marchitto,1,2*† Scott J. Lehman,1,2* Joseph D. Ortiz,3 Jacqueline Flückiger,2‡ Alexander van Geen4 We reconstructed the radiocarbon activity of intermediate waters in the eastern North Pacific over the past 38,000 years. Radiocarbon activity paralleled that of the atmosphere, except during deglaciation, when intermediate-water values fell by more than 300 per mil. Such a large decrease requires a deglacial injection of very old waters from a deep-ocean carbon reservoir that was previously well isolated from the atmosphere. The timing of intermediate-water radiocarbon depletion closely matches that of atmospheric carbon dioxide rise and effectively traces the redistribution of carbon from the deep ocean to the atmosphere during deglaciation. adiocarbon measurements of calendrically dated hermatypic corals (1) and planktonic foraminifera (2, 3) indicate that the radiocarbon activity (D14C) of the atmosphere during the latter part of the last glacial period [~20,000 to 40,000 years before the present (yr B.P.)] ranged from ~300 to 800 per mil (‰) higher than it was during the prenuclear modern era (Fig. 1C). Although reconstructions of Earth’s geomagnetic-field intensity predict higher cosmogenic 14C production rates during the glacial period, production was apparently not high enough to explain the observed atmospheric enrichment (2–5). Rather, a substantial fraction of the atmosphere’s D14C buildup must have been due to decreased uptake of 14C by the deep ocean. This requires a concomitant 14C depletion in a deep-ocean dissolved inorganic C reservoir that was relatively well isolated from the atmosphere. Renewed ventilation of this reservoir could theoretically explain the drop in atmospheric D14C (Fig. 1C) and the rise in atmospheric CO2 (6) across the last deglaciation. Most workers point to the Southern Ocean as a R 1 Department of Geological Sciences, University of Colorado, Boulder, CO 80309, USA. 2Institute of Arctic and Alpine Research, University of Colorado, Boulder, CO 80309, USA. 3 Department of Geology, Kent State University, Kent, OH 44242, USA. 4Lamont-Doherty Earth Observatory, Columbia University, Palisades, NY 10964, USA. *These authors contributed equally to this work. †To whom correspondence should be addressed. E-mail: [email protected] ‡Present address: Environmental Physics, Institute of Biochemistry and Pollutant Dynamics, Eidgenössische Technische Hochschule Zürich, 8092 Zürich, Switzerland. 1456 locus of deglacial CO2 release, based on the similarity between atmospheric CO2 and Antarctic temperature records (6) and on numerous conceptual and numerical models (7–9). If correct, we would expect some signature of the low-14C deepocean C reservoir to be spread to other basins via Antarctic Intermediate Water (AAIW). Here, we report a strong radiocarbon signal of the deglacial release of old C, recorded in an intermediatedepth sediment core from the northern edge of the eastern tropical North Pacific. Intermediate water D14C reconstruction. Marine sediment multi-core/gravity-core/pistoncore triplet from sediment layer MV99-MC19/ GC31/PC08 was raised from a water depth of 705 m on the open margin off the western coast of southern Baja California (23.5°N, 111.6°W) (10). The site is today situated within the regional O2 minimum zone that exists because of a combination of high export production and poor intermediate-water ventilation. Various sediment properties in MC19/GC31/PC08 vary in concert with the so-called Dansgaard-Oeschger (D-O) cycles that characterized the Northern Hemisphere climate during the last glacial period (11). Originally discovered in Greenland ice cores, D-O cycles also exist in a number of lower-latitude locations that were probably teleconnected to the North Atlantic region through the atmosphere (2, 12, 13). Off the coast of Baja California, the sedimentary concentrations of organic C, Cd, Mo, and benthic foraminifera all decreased sharply during D-O stadials (cold periods in Greenland) (11, 14). Together, these proxies are consistent with reduced productivity during stadials, caused 8 JUNE 2007 VOL 316 SCIENCE Supporting Online Material www.sciencemag.org/cgi/content/full/1140429/DC1 Materials and Methods SOM Text Fig. S1 References SOM Programs 25 January 2007; accepted 30 April 2007 Published online 24 May 2007; 10.1126/science.1140429 Include this information when citing this paper. by either decreased coastal upwelling or a deepening of the regional nutricline related to the mean state of the tropical Pacific (11). Diffuse spectral reflectance (DSR) provides a 1-cm resolution stratigraphy for GC31/PC08. After R-mode factor analysis, the third factor of DSR (Fig. 1A) exhibits the strongest correlation to the productivity proxies and to Greenland climate (11). We used this DSR record to apply a calendar-age model to MC19/GC31/PC08, based on correlation to d18O (an air-temperature proxy) in Greenland ice core GISP2 (Greenland Ice Sheet Project 2) (15). Resulting calendar ages were then combined with 50 benthic foraminiferal radiocarbon ages [19 of which were published previously (10)] to calculate age-corrected intermediate-water D14C (16). To evaluate the partitioning of 14C between the atmosphere and the ocean, we compared intermediate-water D14C to that of the atmosphere (Fig. 1C), as reconstructed from tree rings (17), U-Th–dated corals (1, 17), and planktonic foraminifera from Cariaco Basin off Venezuela (3). Calendar ages for Cariaco Basin were originally based on the correlation of lithologic climate proxies to the GISP2 d18O record (2), which has been layercounted with visual and chemical techniques (15). However, Hughen et al. (3) recently demonstrated that the Cariaco Basin 14C calibration yields much better agreement with coral results older than ~22,000 yr B.P. when an alternate age model is used, based on correlation to the U-Th–dated Hulu Cave speleothem d18O record from eastern China (13). Because DSR in GC31/PC08 is more similar to the Greenland isotope record than to the lower-resolution Hulu Cave record, we continued to use the GISP2 correlation but applied simple provisional age adjustments to GISP2 older than 23,400 yr B.P., using four tie points to Hulu Cave (Fig. 1B and fig. S1). We do not suggest that this age model is necessarily superior to the original one (15), but this exercise is necessary for comparing our data to the most recent (and most consistent) atmospheric D14C reconstructions (1, 3, 17). The resulting age model for MC19/GC31/PC08, based on 21 tie points, yields a very constant sedimentation rate (fig. S2) and gives us confidence that our calendar-age assignments for 14C samples between tie points are reliable to within a few hundred years (table S1). Baja California intermediate-water radiocarbon activities are plotted in red in Fig. 1C. The www.sciencemag.org Downloaded from www.sciencemag.org on June 7, 2007 28. 29. 30. 31. 32. 33. modern activity, based on a local seawater measurement of –131‰ at 445 m and the nearest Geochemical Ocean Section Study profile (18), is estimated to be –170‰, in good agreement with the core top value. Comparable offsets from the contemporaneous atmosphere were maintained throughout the Holocene (typically ~100‰ between 0 and 10,000 yr B.P.) and during the latter part of the glacial period (roughly 200‰ between 20,000 and 30,000 yr B.P.) (19). Radiocarbon activities before 30,000 yr B.P. show increased scatter that may be related to the greater influence of slight contaminations on older samples. Overall, it is clear that intermediate waters mainly followed atmospheric D14C over the past 40,000 yr, except during the last deglaciation. Activities dropped sharply just after 18,000 yr B.P., reaching minimum values of ~–180‰ between 15,700 and 14,600 yr B.P., roughly 450‰ lower than that of the contemporaneous atmosphere. For comparison, the lowest D14C values found in the modern ocean (in the North Pacific near 2-km water depth) are depleted by ~240‰, relative to the preindustrial atmosphere (18). A second comparably large depletion event began sometime between 13,500 and 12,900 yr B.P. and ended between 12,100 and 11,600 yr B.P. The magnitude of 14C depletion we reconstructed during these two deglacial events is much too great to attribute to changes in dynamics of the North Pacific thermocline (20) and must instead record a large change in the initial 14C activity of waters advected to the site. Such depleted waters could have been sourced only from the deepest, most isolated regions of the glacial ocean (21). Link to atmospheric history. In Fig. 2, we compare the timing of reconstructed intermediatewater and atmospheric D14C changes with the deglacial record of atmospheric CO2 from the East Antarctic Dome C ice core (6). The latter is shown on a GISP2 layer-counted time scale based on a simple synchronization of CH4 variations between the Dome C and GISP2 ice cores (supporting online material text and table S2). It is immediately apparent that the atmospheric D14C decline and CO2 rise occurred in parallel, with a synchronous, intervening plateau appearing in both records. There is a slight (and still unreconciled) difference in the timing of the start of the deglacial atmospheric D14C decline between the Cariaco Basin (3) and IntCal04 (17) reconstructions, but we take the deglacial onset of the atmospheric CO2 rise and the atmospheric 14 C decline to be essentially synchronous. We now also find that these changes were associated with a prominent decline in D14C of intermediate water in the eastern North Pacific that must record the redistribution of aged C from the deep ocean to the surface. After 14,600 yr B.P., intermediate-water activities rebounded to higher values, coincident with the plateau in both the atmospheric CO2 rise and the atmospheric D14C drop. The leveling of the atmospheric records and the increase in 14C activity of intermediate waters are all indicative of a reduction in the flux of aged C to the upper ocean and atmosphere from below. After ~12,800 yr B.P., the atmospheric CO2 rise and D14C drop resumed, and intermediate-water activities again reached minimum values of ~–180‰, indicating a resumption of C redistribution from the deep ocean to the surface. By 11,500 yr B.P., the large deglacial atmospheric shifts were mostly completed, and intermediate-water activities finally reached modern values. As pointed out by Monnin et al. (6), the deglacial rise of atmospheric CO2 closely followed the rise in East Antarctic temperatures (Fig. 3A), implying that the ocean’s release of C to the atmosphere was associated with changes in the Southern Ocean. Deep convection of the Southern Ocean both ventilates much of the ocean interior and returns to the atmosphere much of the C extracted by photosynthesis in the sunlit surface of the global ocean. During the last Fig. 1. Intermediate-water and atmospheric D14C records. (A) Diffuse spectral reflectance factor 3 from Baja California composite sediment core MV99-GC31/ PC08, plotted versus depth (top axis) (11). Gray lines show tie points to the Greenland record that were used to derive the calendar-age model. (B) d18O of Greenland ice core GISP2 (15) on a provisional revised time scale (black, bottom axis). The new time scale deviates from the original time scale (green) for calendar ages older than 23,400 yr B.P. (vertical line labeled G). The new time scale is based on linear interpolation between point G and three tie points whose ages are derived from U-Th–dated Hulu Cave (vertical lines labeled H) (13). (C) Atmospheric radiocarbon activities based on tree rings, planktonic foraminifera from Cariaco Basin varve-counted sediments, and U-Th–dated corals (dark green) (17); additional recent coral measurements (light blue) (1); and planktonic foraminifera from Cariaco Basin, based on an age model derived from the correlation of sediment reflectance to Hulu Cave (dark blue) (3). Red circles show intermediate-water activities from benthic foraminifera in MC19/GC31/PC08. The yellow circle is an estimate for modern bottom waters at this site. Error bars are based on compounded uncertainties in radiocarbon ages and calendar ages. www.sciencemag.org SCIENCE VOL 316 8 JUNE 2007 Downloaded from www.sciencemag.org on June 7, 2007 RESEARCH ARTICLES 1457 glacial period, density stratification of the Southern Ocean surface and/or extensive sea-ice coverage are suggested to have isolated deep waters from the atmosphere (7–9), permitting the buildup of a larger deep-ocean C reservoir and a consequent drawdown of atmospheric CO2. Sediment pore water chlorinity and d18O measurements, combined with benthic foraminiferal d18O, indicate that deep Southern Ocean waters were the saltiest and densest waters in the glacial ocean (22). Such high salinities point to brine formation beneath sea ice as an important mode of formation. At deglaciation, a progressive renewal of deep convection or upwelling in association with documented sea-ice retreat (23) [and possibly with poleward-shifting westerlies (8)] would have provided for the simultaneous delivery of ocean heat and sequestered C to the atmosphere. This transition occurred in two major steps, beginning with relatively early (~18,000 years ago) and gradual increases in temperature and CO2 that were temporarily interrupted by the Antarctic Cold Reversal (ACR). Major transients in our record of D14C in intermediate-depth waters of the eastern North Pacific conform to this Antarctic schedule, consistent with the redistribution of C from the abyss to the upper ocean and atmosphere in connection with changes in deep convection of the Southern Ocean. d13C provides a tracer that is complementary to D14C, though with a far smaller dynamic range in seawater. During the last glacial period, the deep Southern Ocean contained the ocean’s lowest d13C values, suggesting a local accumulation of remineralized C and/or poor ventilation (24). Spero and Lea (25) argued that during the early part of the last deglaciation, Southern Ocean sea-ice retreat (23), combined with increased deep convection and northward Ekman transport, imparted transient low d13C values to AAIW and Subantarctic Mode Waters and that this signal was recorded by deep-dwelling planktonic foraminifera in various ocean basins, including the eastern tropical North Pacific. These waters should also have carried a low D14C signature, and we suggest that this is the signal we observe off the coast of Baja California. Today, AAIW is barely traceable as a distinct water mass north of the equator in the eastern tropical Pacific (26). We argue that the northward penetration of AAIW during deglaciation was greater than it is today (27) and was at times fed by extremely 14C-depleted waters sourced from the abyss by deep overturning in the Southern Ocean. Sea-ice retreat could have allowed the upwelled deep waters to gain buoyancy from precipitation, converting some fraction of these waters into AAIW without substantial mixing with warmer thermocline waters (28), which would otherwise dampen the D14C signal. There is evidence that vertical stratification of the North Pacific also varied on an Antarctic climate schedule (29), so northern deep waters may have supplemented the supply of aged C to the Baja California site. However, 1458 Fig. 2. Baja California intermediate-water D14C during the last deglaciation (red), compared with atmospheric D14C (dark green, light blue, and dark blue) (1, 3, 17), and atmospheric CO2 from Antarctica Dome C (6) placed on the GISP2 time scale (black), as discussed in the text. Vertical dashed gray lines show the ages of Bølling-Allerød (B-A) and Younger Dryas (YD) boundaries, based on the GISP2 d18O record, and the start of Heinrich event 1 (H1), based on the 231Pa/230Th record from Bermuda Rise (34). The ACR is contemporaneous with the Bølling-Allerød. ppmv, parts per million by volume. the interaction of strong circumpolar winds with bathymetry in the Southern Ocean provides for much more effective vertical pumping (30) than do conditions in the North Pacific, and therefore southern sources probably dominated the D14C changes in our record. North-south teleconnections. Numerous observational and modeling studies indicate an inverse relationship between Antarctic and North Atlantic temperature variations that may be due to altered interhemispheric ocean-heat transport and/or opposing local deep-water formation histories (28, 31–33). Insofar as D14C of the intermediate-depth Pacific provides an inverse proxy for the strength of deep convection in the Southern Ocean, our results provide strong evidence for tight, inverse coupling of deep-water formation between hemispheres. This is clearly demonstrated by the covariation of intermediate-depth Pacific D14C and the formation history of North Atlantic Deep Water (NADW) [or its glacial analog, Glacial North Atlantic Intermediate Water (GNAIW)], based on measurements of 231Pa/230Th from sediments in the western North Atlantic (34) (Fig. 3C). The near-cessation of 231Pa export beginning just after 18,000 yr B.P. records a collapse of GNAIW that has been linked to a massive discharge of glacial ice and fresh water to the North Atlantic, known as Heinrich event 1. After a recovery during the Bølling-Allerød warm phase, another marked weakening of NADW/ GNAIW is documented during the Younger 8 JUNE 2007 VOL 316 SCIENCE Dryas cold period, probably also triggered by a discharge of glacial meltwater (34). Both periods of NADW/GNAIW reduction were times of intermediate-depth Pacific D14C decline and atmospheric CO2 rise. It is often difficult to identify triggers in a tightly coupled system, but the relationships described above suggest that the Antarctic climate schedule may have been paced by ice sheet and meltwater forcing around the North Atlantic. Support for this conclusion comes from our observation that although major inflections in the D14C and 231Pa/230Th records in Fig. 3 were almost exactly synchronous, the large D14C decrease (i.e., Southern Ocean ventilation increase) during Heinrich event 1 occurred more gradually than did the associated decrease in GNAIW export. This relationship is consistent with relatively slow circum-Antarctic warming due to anomalous ocean-heat transport (33), leading to sea-ice retreat (23) [and possibly poleward-shifting westerlies (8)] and, consequently, a progressive increase in deep overturning of the Southern Ocean. Alternatively, Southern Ocean overturning may have been instigated by NADW/GNAIW reductions through the requirement (35) that global rates of deep-water formation balance global deep upwelling, which is forced mainly by winds and tides. The abrupt rise in atmospheric D14C at the start of the Younger Dryas [+80‰ in just 180 years (36)] (Fig. 2) may record the time elapsed before the Southern Ocean could begin respond- www.sciencemag.org Downloaded from www.sciencemag.org on June 7, 2007 RESEARCH ARTICLES Fig. 3. Southern and northern ocean-atmosphere changes during the last deglaciation, compared with intermediatewater D14C. (A) Atmospheric CO2 (black) and ice core deuterium (dD) temperature proxy (light blue) from Antarctica Dome C (6), placed on the GISP2 time scale. (B) Baja California intermediate-water D14C. (C) Inverted decaycorrected excess 231Pa/230Th in Bermuda Rise sediments, based on two methods to calculate excess (green and purple) (34). The horizontal dashed line shows the watercolumn production ratio for these isotopes (0.093); lower values are primarily due to Pa export by vigorous NADW. Vertical dashed lines show the ages of climatic boundaries, as in Fig. 2. Error bars are based on compound uncertainties in radiocarbon ages and calendar ages. ing to reduced NADW formation, leading to the brief absence of deep-ocean sinks for 14C (32). Recent work shows that deep North Atlantic radiocarbon activities also increased abruptly during the Bølling-Allerød because of renewed formation of NADW and that they temporarily decreased during the Younger Dryas, when NADW formation was again briefly restricted (37, 38). In light of our new record, this pattern of deep-ocean change was probably limited to the North Atlantic (arising from deep circulation changes within the basin), whereas the intermediate-depth North Pacific record tracks the overall redistribution of C from the deep ocean to the atmosphere. Implications for the deep C reservoir. Finally, our results bear on recent questions concerning reconstructed rates of atmospheric D14C decline during deglaciation and implied 14C aging of glacial deep waters. Although subject to uncertainties (19), decay projection of our first deglacial D14C minimum back to the surface ocean (39) gives an apparent ventilation age of ~4000 years, implying that the inferred deep Southern Ocean source waters were at least that old. This is broadly consistent with the minimum deep-ocean age estimated from the atmospheric record, assuming that the old reservoir filled half of the ocean’s volume (40). Ages of up to 5000 years have been reported for glacial deep waters near New Zealand (41), but more northerly sites in the Pacific show little difference from today, at least at depths shallower than ~2 km (40). We infer that the greatest 14C depletion of the glacial deep ocean was probably concentrated in the Southern Ocean region (and deepest Pacific), coincident with the highest densities (22) and lowest d13C values (24). References and Notes 1. R. G. Fairbanks et al., Quat. Sci. Rev. 24, 1781 (2005). 2. K. Hughen et al., Science 303, 202 (2004). 3. K. Hughen, J. Southon, S. J. Lehman, C. Bertrand, J. Turnbull, Quat. Sci. Rev. 25, 3216 (2006). 4. C. Laj et al., Earth Planet. Sci. Lett. 200, 177 (2002). 5. R. Muscheler et al., Earth Planet. Sci. Lett. 219, 325 (2004). 6. E. Monnin et al., Science 291, 112 (2001). 7. D. M. Sigman, E. A. Boyle, Nature 407, 859 (2000). 8. J. R. Toggweiler, J. L. Russell, S. R. Carson, Paleoceanography 21, PA2005 (2006). 9. B. B. Stephens, R. F. Keeling, Nature 404, 171 (2000). 10. A. van Geen et al., Paleoceanography 18, 1098 (2003). 11. J. D. Ortiz et al., Geology 32, 521 (2004). 12. R. J. Behl, J. P. Kennett, Nature 379, 243 (1996). 13. Y. J. Wang et al., Science 294, 2345 (2001). 14. W. E. Dean, Y. Zheng, J. D. Ortiz, A. van Geen, Paleoceanography 21, PA4209 (2006). www.sciencemag.org SCIENCE VOL 316 15. P. M. Grootes, M. Stuiver, J. Geophys. Res. 102, 26455 (1997). 16. Materials and methods are available as supporting material on Science Online. In the supporting material, we also present our D14C values calculated with the original GISP2 chronology (15) and compare them to the original Cariaco record (2), based on correlation to GISP2 (fig. S3). 17. P. J. Reimer et al., Radiocarbon 46, 1029 (2004). 18. G. Ostlund, H. Craig, W. S. Broecker, D. Spencer, GEOSECS Atlantic, Pacific and Indian Ocean Expeditions, vol. 7, Shorebased Data and Graphics (U.S. Government Printing Office, Washington, DC, 1987). 19. Throughout most of this paper, we discuss our D14C values with respect to the contemporaneous atmosphere, rather than calculating ventilation ages by decay projection (39) because of large uncertainties in initial surface-ocean reservoir ages and the possibility of multiple intermediate- and deep-water sources with different ventilation histories. 20. U. Mikolajewicz, T. J. Crowley, A. Schiller, R. Voss, Nature 387, 384 (1997). 21. The deglacial injection of low-14C intermediate waters along the Baja California margin did not noticeably overprint the DSR or other O2-sensitive productivity proxies recorded in GC31/PC08. Although D14C and O2 are broadly correlated in the modern ocean, they can also be decoupled (18) through air-sea exchange. Hence, deglacial intermediate waters were very depleted in D14C but apparently not drastically depleted in O2. 22. J. F. Adkins, K. McIntyre, D. P. Schrag, Science 298, 1769 (2002). 23. R. Gersonde, X. Crosta, A. Abelmann, L. Armand, Quat. Sci. Rev. 24, 869 (2005). 24. U. S. Ninnemann, C. D. Charles, Earth Planet. Sci. Lett. 201, 383 (2002). 25. H. J. Spero, D. W. Lea, Science 296, 522 (2002). 26. P. C. Fiedler, L. D. Talley, Prog. Oceanogr. 69, 143 (2006). 27. S. Schulte, F. Rostek, E. Bard, J. Rullkötter, O. Marchal, Earth Planet. Sci. Lett. 173, 205 (1999). 28. R. F. Keeling, B. B. Stephens, Paleoceanography 16, 112 (2001). 29. S. L. Jaccard et al., Science 308, 1003 (2005). 30. J. R. Toggweiler, B. Samuels, Deep-Sea Res. 42, 477 (1995). 31. T. J. Crowley, Paleoceanography 7, 489 (1992). 32. W. S. Broecker, Paleoceanography 13, 119 (1998). 33. R. Knutti, J. Flückiger, T. F. Stocker, A. Timmermann, Nature 430, 851 (2004). 34. J. F. McManus, R. Francois, J.-M. Gherardi, L. D. Keigwin, S. Brown-Leger, Nature 428, 834 (2004). 35. W. Munk, C. Wunsch, Deep-Sea Res. 45, 1977 (1998). 36. K. A. Hughen, J. R. Southon, S. J. Lehman, J. T. Overpeck, Science 290, 1951 (2000). 37. L. C. Skinner, N. J. Shackleton, Paleoceanography 19, PA2005 (2004). 38. L. F. Robinson et al., Science 310, 1469 (2005). 39. J. F. Adkins, E. A. Boyle, Paleoceanography 12, 337 (1997). 40. W. Broecker et al., Science 306, 1169 (2004). 41. E. L. Sikes, C. R. Samson, T. P. Guilderson, W. R. Howard, Nature 405, 555 (2000). 42. We thank D. Lopez, J. Turnbull, and C. Wolak for laboratory assistance; J. Southon for accelerator mass spectrometry analyses; A. Pearson for providing an unpublished seawater D14C measurement; and T. Blunier for providing Dome C EDC 3 time scale and for valuable discussions on synchronization of the different ice cores. This manuscript was improved by comments from R. Keeling and two anonymous reviewers. Support was provided by NSF grants OCE-9809026 and OCE-0214221. Downloaded from www.sciencemag.org on June 7, 2007 RESEARCH ARTICLES Supporting Online Material www.sciencemag.org/cgi/content/full/1138679/DC1 Materials and Methods SOM Text Figs. S1 to S3 Tables S1 and S2 References 11 December 2006; accepted 27 April 2007 Published online 10 May 2007; 10.1126/science.1138679 Include this information when citing this paper. 8 JUNE 2007 1459 www.sciencemag.org/cgi/content/full/1138679/DC1 Supporting Online Material for Marine Radiocarbon Evidence for the Mechanism of Deglacial Atmospheric CO2 Rise T. M. Marchitto,* S. J. Lehman, J. D. Ortiz, J. Flückiger, A. van Geen *To whom correspondence should be addressed. E-mail: [email protected] Published 10 May 2007 on Science Express DOI: 10.1126/science.1138679 This PDF file includes: Materials and Methods SOM Text Figs. S1 to S3 Tables S1 and S2 References Supporting Online Material Methods Bulk sediment samples were washed using a sodium hexametaphosphate solution and picked for benthic foraminifera using the >250 µm size fraction, supplemented with 150250 µm specimens where necessary. Toward the end of this study, picked foraminifera were also briefly sonicated in methanol (Table S1). Foraminiferal samples were hydrolyzed with H3PO4 and reduced to graphite using an Fe catalyst in the presence of H2. The 19 published radiocarbon samples (S1) were graphitized at the National Ocean Sciences AMS Facility in Woods Hole (NOSAMS), while the 31 new samples were graphitized at the INSTAAR Laboratory for AMS Radiocarbon Preparation and Research at the University of Colorado, Boulder (NSRL). All published radiocarbon data (S1) were analyzed at NOSAMS, while all new data were analyzed at the Keck Carbon Cycle AMS Facility at the University of California, Irvine (KCCAMS). δ13C corrections were based on off-line measurement for NOSAMS results, and on measurement within the AMS for KCCAMS results. KCCAMS δ13C values are not suitable for paleoceanographic interpretation, and are therefore not reported. Age-corrected ∆14C calculations were based on the conventions of Stuiver and Polach (S2). ∆14C error bars were calculated by compounding radiocarbon age errors with estimated calendar age errors listed in Table S1. Calendar age errors depend solely on the estimated precision of our picks with respect to the GISP2-Hulu chronology, and not on any errors in the GISP2-Hulu chronology itself. The latter would affect both our record and the Cariaco Basin atmospheric ∆14C record similarly. Hulu Cave adjustments to GISP2 age model In order to compare PC08 results with the most recent and most consistent atmospheric ∆14C reconstructions (S3-S5), the correlation between PC08 reflectance and GISP2 δ18O (S6) (used to derive calendar ages for calculating PC08 ∆14C) requires that the GISP2 chronology be consistent with the Hulu Cave chronology (S7), as outlined in the main text. Younger than ~20-25 kyr BP the Hulu Cave and GISP2 chronologies appear to be consistent with each other, so no corrections need to be applied to the more recent parts of the GISP2 age model (Fig. S1). GISP2 also agrees with the latest NGRIP chronology (GICC05) to within 250 yr over this interval (S8). We adjusted the GISP2 chronology older than 23.4 kyr BP (the start of Interstadial 2) by linearly interpolating between three Hulu Cave tie-points corresponding to major interstadial warmings in Greenland. The resulting GISP2 age model appears to agree with Hulu Cave to within a few hundred years throughout the past 50 kyr (Fig. S1). The resulting age-depth relationship for PC08 is shown in Fig. S2A. Methane synchronization of Dome C and GISP2 As outlined above and in the main text, the GISP2, Hulu Cave, and coral chronologies are similar over the last ~20-25 kyr. However, these chronologies differ from the glaciological age model on which the EPICA Dome C CO2 and δD records of Monnin et al. (S9) were originally presented. To obtain a timescale that is consistent with the various ∆14C reconstructions, we have used CH4 records from the Dome C, GISP2, and GRIP ice cores (S9-S11) to derive a GISP2 synchronized gas age scale for Dome C. Seven tie-points (Table S2) were defined based on the comparison of the Dome C CH4 record (on the EDC3 time scale (S12, S13)) and the GISP2 and GRIP CH4 records (on the GISP2 time scale (S10)). Between the tie-points the EDC3 gas age scale was linearly interpolated to GISP2 ages. The associated ice age scale for Dome C was derived from the synchronized gas age scale described above and the delta ages (gas age to ice age differences in the ice core) taken from the EDC3 time scale (S12, S13). Sedimentary artifacts The two deglacial intervals of depleted radiocarbon activity that we report appear, in part, as large age plateaus in the conventional radiocarbon age:depth relationship (Table S1). During the first (earlier) event, radiocarbon age is nearly constant over 90 cm of core depth, and during the second event, radiocarbon age is nearly constant over at least 45 cm. Near-instantaneous massive inputs of sediment, namely slumps, could theoretically produce age plateaus. However, such events would necessarily produce large kinked offsets in the calendar age:depth relationship, which are clearly not observed (Fig. S2). Additionally, slumps are much more likely to produce age reversals than plateaus. Radiocarbon age plateaus could also theoretically be created during intervals of very intense bioturbation, such that age is homogenized over some depth interval. However, it is unreasonable to postulate a complete, simultaneous bioturbational homogenization of the upper 90 cm (or 45 cm) of the sediment column, especially in this low-O2 environment. The entire MC19/GC31/PC08 record is dominated by low-O2 benthic foraminiferal taxa (mainly Bolivina and Uvigerina), and sedimentary Mo remains enriched by ~4-8 times crustal values during deglaciation, indicative of shallow pore water sulfide and suboxic bottom waters (S14). Finally, very intense bioturbation would homogenize other paleoclimate proxies in this core. Notably, our second age plateau crosses the Allerød-Younger Dryas boundary in GC31/PC08, which is marked by a sharp reflectance change that is inferred to record a shift in surface ocean productivity (S15). We therefore conclude decisively that the 14C age-depth patterns in our record reflect changes in intermediate water 14C activity and not sedimentary processes. Alternate PC08 age model We have also calculated Baja California ∆14C based on correlation to GISP2 using the original layer-counted age model of Meese et al. (S16) (Fig. S3). This record only differs from our preferred GISP2-Hulu age model prior to 23.4 kyr BP. In Fig. S3 we compare this alternate record to the original Cariaco Basin ∆14C record based on correlation to GISP2 (S17). It is clear that our conclusions are not affected by the choice of age model, though the agreement between Cariaco Basin and coral ∆14C is better in Fig. 1 than in Fig. S3 (S5). The PC08 age-depth relationship for the alternate age model is shown in Fig. S2B. 2 Fig. S1. Derivation of GISP2-Hulu Cave composite age model. (A) Diffuse spectral reflectance factor 3 from Baja California composite sediment core MV99-GC31/PC08, plotted versus depth (S15). Gray lines show tie-points to Greenland record used to derive calendar age model. (B) δ18O of Greenland ice core GISP2 (S6) on provisional revised timescale (black). New timescale deviates from original timescale (green) older than 23.4 kyr BP (line labeled G). New timescale is based on linear interpolation between point G and three tie-points whose ages are derived from U-Th-dated Hulu Cave (lines labeled H). (C) Hulu Cave δ18O from three different stalagmites (S7). Gray lines indicate tiepoints used to adjust GISP2 chronology. 3 Fig. S2. (A) Age-depth relationship for MV99-MC19/GC31/PC08 based on correlation to GISP2 δ18O with Hulu Cave age adjustment. Red points are tie-points and gray points represent radiocarbon measurements. The unusually constant sedimentation rate allows for relatively precise calendar age assignments between tie-points. (B) Same as (A) except using alternate age model as in Fig. S3. 4 Fig. S3. Baja California ∆14C record based on alternate age model. (A) Diffuse spectral reflectance factor 3 from Baja California composite sediment core MV99-GC31/PC08, plotted versus depth (top axis) (S15). Gray lines show tie-points to Greenland record used to derive alternate calendar age model. (B) δ18O of Greenland ice core GISP2 on original timescale (bottom axis) (S6, S16). (C) Atmospheric radiocarbon activities based on tree rings, planktonic foraminifera from Cariaco Basin varve-counted sediments, and U-Thdated corals (dark green) (S3); additional recent coral measurements (cyan) (S4); and planktonic foraminifera from Cariaco Basin using age model derived from reflectance correlation to GISP2 (blue) (S17). Red points show intermediate water activities from benthic foraminifera in MC19/GC31/PC08. Yellow point is estimate for modern bottom waters at this site. 5 Sample MC19-10 cm MC19-25 cm GC31-3-2 cm MC19-40 cm GC31-3-22.5 cm GC31-3-50.5 cm GC31-3-70 cm GC31-3-100.5 cm GC31-3-123 cm GC31-2-0.5 cm GC31-2-50.5 cm GC31-2-100.5 cm GC31-1-8.5 cm PC08-9-45 cm PC08-9-60 cm GC31-1-32.5 cm PC08-9-85 cm GC31-1-58.5 cm PC08-9-110 cm PC08-9-130 cm GC31-1-94.5 cm PC08-9-141 cm PC08-9-150 cm PC08-8-6.5 cm PC08-8-16.5 cm PC08-8-25.5 cm PC08-8-35 cm Taxa mixed benthics mixed benthics mixed benthics mixed benthics mixed benthics Bolivina spp. mixed benthics Bolivina spp. mixed benthics Bolivina spp. Bolivina spp. Bolivina spp. Bolivina spp. mixed benthics mixed benthics mixed benthics mixed benthics Bolivina spp. mixed benthics mixed benthics Bolivina spp. Uvigerina spp. mixed benthics Uvigerina spp. Uvigerina spp. Uvigerina spp. mixed benthics Composite depth (m) 0.100 0.250 0.270 0.400 0.475 0.755 0.950 1.255 1.480 1.755 2.255 2.755 3.255 3.260 3.410 3.495 3.660 3.755 3.910 4.110 4.115 4.220 4.310 4.375 4.475 4.565 4.660 C age (kyr BP) 1.720 2.050 2.050 2.320 2.230 3.030 3.690 3.840 5.130 5.810 7.190 8.980 10.050 10.460 10.845 11.600 13.380 13.500 13.530 13.420 13.650 14.285 13.370 14.485 15.755 15.850 16.505 14 Table S1. Core MC19/GC31/PC08 radiocarbon data. C age error (kyr) 0.030 0.035 0.030 0.035 0.035 0.040 0.045 0.050 0.060 0.040 0.050 0.060 0.410 0.030 0.030 0.070 0.035 0.070 0.030 0.025 0.150 0.035 0.030 0.035 0.040 0.040 0.040 14 Calendar age (kyr BP) 0.32 0.81 0.87 1.29 1.54 2.44 3.08 4.07 4.79 5.69 7.31 9.01 10.79 10.81 11.32 11.61 12.12 12.41 12.88 13.57 13.58 13.94 14.20 14.37 14.65 14.96 15.30 Calendar age error (kyr) 0.1 0.2 0.3 0.4 0.4 0.4 0.4 0.4 0.4 0.4 0.3 0.3 0.2 0.2 0.2 0.1 0.2 0.2 0.1 0.1 0.3 0.1 0.2 0.2 0.1 0.2 0.3 ∆14C (per mil) -161 -146 -139 -124 -88 -78 -84 14 -57 -35 -11 -27 56 6 20 -39 -181 -164 -118 -29 -54 -88 55 -62 -172 -150 -184 -1.43 -1.49 -1.49 δ13C (per mil) -0.61 -0.69 -0.86 -0.56 -0.91 nm -0.60 nm -0.53 nm nm -0.90 -1.15 Accession # OS-33198 OS-33199 OS-33201 OS-33200 OS-25612 OS-22946 OS-33202 OS-22947 OS-33203 OS-22948 OS-22949 OS-22955 OS-23513 CURL-8750 CURL-8752 OS-25611 CURL-8751 OS-22956 CURL-8444 CURL-8445 OS-22957 CURL-8746 CURL-8446 CURL-8721 CURL-8726 CURL-8720 CURL-8447 Son? no no no no no no no no no no no no no yes yes no yes no no no no yes no yes yes yes no 6 Reference 1 1 1 1 1 1 1 1 1 1 1 1 1 this study this study 1 this study 1 this study this study 1 this study this study this study this study this study this study 7 PC08-8-45.5 cm Uvigerina spp. 4.765 16.785 0.045 15.67 0.3 -176 CURL-8724 yes this study PC08-8-55.5 cm Uvigerina spp. 4.865 16.665 0.040 16.02 0.3 -127 CURL-8729 yes this study PC08-8-65.5 cm Uvigerina spp. 4.965 16.425 0.045 16.37 0.3 -62 CURL-8744 yes this study PC08-8-75.5 cm Uvigerina spp. 5.065 16.390 0.040 16.73 0.3 -17 CURL-8742 yes this study PC08-8-85.5 cm Uvigerina spp. 5.165 16.425 0.050 17.08 0.3 22 CURL-8743 yes this study PC08-8-95.5 cm Uvigerina spp. 5.265 16.185 0.045 17.43 0.3 98 CURL-8749 yes this study PC08-8-105.5 cm Uvigerina spp. 5.365 16.235 0.040 17.78 0.3 139 CURL-8748 yes this study PC08-8-116 cm Uvigerina spp. 5.470 16.810 0.040 18.12 0.3 105 CURL-8745 yes this study PC08-8-125 cm mixed benthics 5.560 16.680 0.040 18.41 0.3 162 CURL-8448 no this study PC08-7-60.5 cm Uvigerina spp. 6.415 19.720 0.060 21.14 0.2 108 CURL-8728 yes this study PC08-7-80.5 cm Uvigerina spp. 6.615 19.640 0.060 21.77 0.3 208 CURL-8722 yes this study PC08-7-99.75 cm mixed benthics 6.808 19.650 0.080 22.39 0.3 299 -0.81 OS-33204 no 1 PC08-7-132.5 cm Uvigerina spp. 7.135 21.170 0.070 23.43 0.1 220 CURL-8727 yes this study PC08-6-20.5 cm Uvigerina spp. 7.585 22.260 0.070 25.15 0.3 311 CURL-8725 yes this study PC08-6-118 cm mixed benthics 8.560 25.500 0.170 29.00 0.2 396 -0.57 OS-33205 no 1 PC08-5-25 cm mixed benthics 9.010 27.200 0.130 30.73 0.2 394 CURL-8449 no this study PC08-5-95 cm mixed benthics 9.710 29.230 0.160 33.11 0.2 443 CURL-8450 no this study PC08-5-130 cm Uvigerina spp. 10.060 30.830 0.170 34.55 0.2 407 CURL-7188 no this study PC08-5-145 cm Uvigerina spp. 10.210 30.460 0.190 35.11 0.2 577 CURL-7189 no this study PC08-5-145 cm Bolivina spp. 10.210 31.370 0.180 35.11 0.2 408 CURL-7192 no this study PC08-4-15 cm mixed benthics 10.410 31.200 0.280 35.89 0.2 582 -0.72 OS-33206 no 1 PC08-4-15 cm Uvigerina spp. 10.410 31.750 0.190 35.89 0.2 477 CURL-7187 no this study PC08-4-65 cm Bolivina spp. 10.910 34.400 0.260 37.92 0.2 356 CURL-7193 no this study Composite depth is relative to the top of MC19, as in (S1). Calendar age errors are based on estimated precision of tie-points and distance of samples from tiepoints. δ13C data are only reported for NOSAMS measurements; ‘nm’ indicates not measured. Column labeled ‘Son?’ indicates whether or not radiocarbon samples were sonicated in methanol prior to graphitization. Table S2. Tie-points between the EPICA Dome C EDC3 and GISP2 time scales. EDC3 gas age (yr BP) 0 8200 11400 12400 14110 16700 21860 GISP2 gas age (yr BP) 0 8200 11620 12750 14810 17600 22950 estimated uncertainty (yr) 0 25 25 100 150 200 300 event description 8.2 ka BP event YD termination Onset of YD Onset of B-A First deglacial CH4 rise CH4 peak in GRIP and Dome C 8 SOM References S1. S2. S3. S4. S5. S6. S7. S8. S9. S10. S11. S12. S13. S14. S15. S16. S17. A. van Geen et al., Paleoceanography 18, doi:10.1029/2003PA000911 (2003). M. Stuiver, H. A. Polach, Radiocarbon 19, 355 (1977). P. J. Reimer, et al., Radiocarbon 46, 1029 (2004). R. G. Fairbanks et al., Quaternary Science Reviews 24, 1781 (2005). K. Hughen, J. Southon, S. J. Lehman, C. Bertrand, J. Turnbull, Quaternary Science Reviews, 25, 3216 (2006). P. M. Grootes, M. Stuiver, Journal of Geophysical Research 102, 26455 (1997). Y. J. Wang et al., Science 294, 2345 (2001). A. Svensson et al., Quaternary Science Reviews, 25, 3258 (2006). E. Monnin et al., Science 291, 112 (2001). T. Blunier, E. Brook, Science 291, 109 (2001). J. Flückiger et al., Global Biogeochemical Cycles 16, 1010 (2002). F. Parrenin et al., Climate of the Past Discussions 3, 19 (2007). L. Loulergue et al., Climate of the Past Discussions 3, 425 (2007). W. E. Dean, Y. Zheng, J. D. Ortiz, A. van Geen, Paleoceanography 21, doi:10.1029/2005PA001239 (2006). J. D. Ortiz et al., Geology 32, 521 (2004). D. A. Meese et al., Journal of Geophysical Research 102, 26411 (1997). K. Hughen et al., Science 303, 202 (2004). 9 PERSPECTIVES further work on the stress accumulation and dissipation in the lithosphere during subduction is necessary to understand the faulting mechanism causing seismicity in double WBZ or even triple WBZ, as proposed for some regions beneath Japan (6). Brudzinski et al. show that, as the number of seismological stations and the availability of digital seismic traces increases, the global earthquake catalog will become accurate enough to delineate the fine structure of seismicity (on the order of a few kilometers) on a global scale. This accuracy will increase in the near future as a result of large seismological observation initiatives like EarthScope in the United States or the NERIES program (Network of Research Infrastructures for European Seismology) in Europe, which will make more high-quality digital data readily available for seismologists worldwide. References 1. M. R. Brudzinski, C. H. Thurber, B. R. Hacker, E. R. Engdahl, Science 316, 1472 (2007). 2. A. Hasegawa, N. Umino, A. Takagi, Tectonophysics 47, 43 (1978). 3. A. Rietbrock, F. Waldhauser, Geophys. Res. Lett. 31, 10.1029/2004GL019610 (2004). 4. S. Kirby, Rev. Geophys. 33, 287 (1995). 5. B. R. Hacker, G. A. Abers, S. M. Peacock, J. Geophys. Res. 108, 2029 (2003). 6. T. Igarashi, T. Matsuzawa, N. Umino, A. Hasegawa, J. Geophys. Res. 106, 2177 (2001). 10.1126/science.1141921 ATMOSPHERE Results from a sediment core provide insights into ocean circulation changes during the last deglaciation. Deglaciation Mysteries Ralph F. Keeling etween 19,000 and 11,000 waters, which is a measure of Glacial state SOUTH NORTH Cap 0m years ago, the Earth emthe deep-water ventilation rate. Core site erged from the last glacial This technique, applied to period. During this deglaciation, numerous sediment cores, has –2800 m Poorly ventilated deep water the carbon dioxide (CO2) concenthus far mainly yielded the unre14 Low C tration in the atmosphere rose from markable result that the ventila–4000 m 180 to 265 parts per million (ppm). tion rate of the glacial ocean was Over the same period, the radiovery similar to that of today’s CO2 release Early deglacial state 0m carbon content of the CO2 fell by ocean, at least down to a depth Low 14C intermediate water ~35%. A simple but unproven exof ~2800 m (2). Thus, if there planation for both changes is an was a major change in ocean –2800 m increase in the rate at which the ventilation during deglaciation, Poorly ventilated deep water ocean’s subsurface waters were this change must have occurred –4000 m renewed by exchange with aerated in ocean waters below that depth. surface waters—a process known However, despite some tanas ventilation. A ventilation in- Deep-water ventilation. This cross section of the Pacific Ocean shows how poorly talizing results (3), no general crease could increase atmospheric ventilated water may have been delivered to intermediate depths during deglacia- picture has emerged for waters tion, as suggested by Marchitto et al. (Top) Ventilation of the deep ocean by sinkCO2 concentration by releasing below 2800 m, because of mething around Antarctica was partially suppressed by a cap formed by sea ice or a layer excess CO2 that had accumulated of low-salinity water. (Bottom) This cap was removed during early deglacial warm- odological difficulties related in subsurface waters by the decom- ing, exposing upwelled deep waters to the atmosphere, releasing radiocarbon- to the low sedimentation rates position of sinking detritus. On depleted CO . The density of the poorly ventilated waters was reduced by freshen- that typically characterize cores 2 page 1456 in this issue, Marchitto ing and warming. With reduced density, the water could spread widely at interme- from these depths. et al. (1) bolster the case for such diate depths, displacing waters of similar density. Marchitto et al. focus on a a ventilation increase and offer insediment core recently hauled of dissolved carbon. Because of radioactive up off Baja California from a depth of 700 m, sight into how the increase may have occurred. To track changes in past ventilation, most decay, waters that are more isolated from the seemingly too shallow for studying deep venresearchers have turned to measurements of atmosphere (that is, more poorly ventilated) tilation. The core contains bands correspondthe radiocarbon (14C) content of shells of have lower 14C/12C ratios, as do the shells that ing to a set of millennial climate oscillations foraminifera, a ubiquitous marine microor- grow in these waters. The radiocarbon age of a first discovered in ice cores from Greenland, ganism. Radiocarbon is produced naturally in fossil shell therefore reflects the age of the allowing absolute dates to be fixed within the the upper atmosphere by cosmic rays and shell plus the age of the water in which it lived. core. Using these dates, the authors correct the spreads through the ocean as part of the pool By subtracting the radiocarbon ages of sur- 14C/12C ratios of benthic foraminifera for face-dwelling (planktonic) and bottom- radioactive decay, thus establishing the origidwelling (benthic) foraminifera, picked from nal 14C/12C ratio of the water in which the The author is at the Scripps Institution of Oceanography, the same layer of a sediment core, it is possible foraminifera lived. The technique does not University of California, San Diego, CA 92093, USA. to estimate age difference of surface and deep require 14C/12C measurements on planktonic E-mail: [email protected] B 1440 8 JUNE 2007 VOL 316 SCIENCE Published by AAAS www.sciencemag.org Downloaded from www.sciencemag.org on June 7, 2007 Brudzinski et al. have now found a direct correlation whereby older oceanic plates show a greater distance between regions of seismicity. They conclude, based on thermalpetrological models developed by Hacker et al. (5), that dehydration of the mineral antigorite is responsible for the seismicity in the lower layer of double WBZs. Although the detailed geophysical explanation presented by Brudzinski et al. for the double seismic zone might be debatable, they observe that double WBZs are the rule and not the exception during subduction of oceanic lithosphere. This provides an important new constraint for all models developed to explain the occurrence of WBZ seismicity. However, foraminifera, which are subject to potentially large systematic errors. The results show two periods during deglaciation when the bottom water at their site had unexpectedly low 14C/12C ratios. The water was so old that it must have been delivered to the site by upwelling from greater depth, presumably from below 2800 m. The oldest waters found by Marchitto et al. have an age of ~4000 years. For comparison, the oldest waters in the modern ocean have an age of ~2300 years. The study provides the strongest evidence to date that the glacial ocean contained some very poorly ventilated water somewhere in its depths. The low-14C periods coincide with the periods when atmospheric radiocarbon decreased and atmospheric CO2 increased most rapidly during deglaciation. The results are thus a convincing fingerprint of a process that flushed excess carbon from an isolated deep reservoir toward the surface, thereby driving the atmospheric changes. Today, waters below 2800 m are ventilated by two routes. One involves the sinking of aerated surface waters in the North Atlantic, the other sinking of such waters near Antarctica. During the last glacial period, both routes probably weakened, with the southern route possibly influenced by sea ice or surface freshening (see the figure, top panel). During glacial times, the deep ocean would thus have been less ventilated than it is today. But how could low-14C waters get to Marchitto et al.’s core site during deglaciation? Much of the upwelling of deep water occurs today around Antarctica, resulting in the formation of Antarctic Intermediate Water, a lowsalinity water mass that spreads northward at intermediate depths. Marchitto et al. hypothesize that a similar process occurred during deglaciation, allowing upwelled water to spread northward to their site (see the figure, bottom panel). However, the evidence for this southern pathway is circumstantial. The results help to reconcile the reconstructed trends in atmospheric radiocarbon with the estimated trends in the production of radiocarbon by cosmic rays—a comparison that seems to demand an increase in ocean ventilation during deglaciation (4). They support theories that attribute the bulk of the glacial-interglacial CO2 change to changes in ocean ventilation (5, 6). The study also provides support for a theory for how the glacial ocean differed from today’s ocean as a result of the cooling of deep waters to nearly the freezing point. Cooling to this extent is expected to allow the salty brine that is released during sea ice formation to accumulate more easily in the deep ocean (7). This idea is supported by sediment pore-water studies (8). By blocking the input of fresh water from precipitation, sea ice could also reduce the conversion of upwelled deep water into low-salinity Antarctic Intermediate Water (7). A strengthening of intermediate-water formation during deglaciation is consistent with a breakdown of this state caused by warming. The study nevertheless leaves the skeptics MOLECULAR BIOLOGY with arrows in their quiver. Marchitto et al.’s low-14C waters are so old that they start to stretch credibility, especially considering that the deep reservoir from which the water was drawn must have been even older. (This follows because some mixing with younger water would unavoidably have occurred during upwelling and transit to the site.) How could prior studies have overlooked deep waters this old? If Marchitto et al.’s interpretation is correct, evidence for old water at intermediate depths should be present throughout the South Pacific in sediments of the appropriate age and depth. If subsequent work supports the findings, we may look back at this study as a key turning point in the quest to understand glacial and interglacial CO2 changes. References 1. T. M. Marchitto, S. J. Lehman, J. D. Ortiz, J. Flückiger, A. van Geen, Science 316, 1456 (2007); published online 10 May 2007 (10.1126/science. 1138679). 2. W. Broecker et al., Paleoceanography 22, PA2206 (2007). 3. L. D. Keigwin, S. J. Lehman, M. S. Cook, Eos 87, PP44A07 (2006). 4. K. Hughen et al., Science 303, 202 (2004). 5. J. R. Toggweiler, J. L. Russell, S. R. Carson, Paleoceanography 21, PA2005 (2006). 6. B. B. Stephens, R. F. Keeling, Nature 404, 171 (2000). 7. R. F. Keeling, B. B. Stephens, Paleoceanography 16, 112 330 (2001). 8. J. F. Adkins, K. McIntyre, D. P. Schrag, Science 298, 1769 (2002). Published online 10 May 2007; 10.1126/science.1142326. Include this information when citing this paper. An advance in DNA sequencing is a crucial component of a rapid, precise, and relatively inexpensive way to identify transcription factor binding sites at a whole-genome level. Site-Seeing by Sequencing Downloaded from www.sciencemag.org on June 7, 2007 PERSPECTIVES Stanley Fields very few years, a new technology comes along that dramatically changes how fundamental questions in biology are addressed. The impact of the technology is not always appreciated at first—when it is used only by those involved in its development—but becomes clear once the technology begins to spread to the broader scientific community. A well-known example is the DNA microarray, which became widely available to biologists about a decade ago and has since been applied to an ever-expanding set of questions such as E The author is in the Departments of Genome Sciences and Medicine, Howard Hughes Medical Institute, University of Washington, Seattle, WA 98195–5065, USA. E-mail: [email protected] determining the profile of genes expressed in a specific cell type. Now it is ultrahigh-throughput DNA sequencing that is making the transition from development to widespread use. Johnson and colleagues are in the vanguard of this movement. On page 1497 of this issue (1), they report that an advanced DNA sequencing technology (from Solexa/Illumina) can be used to identify all the locations in the human genome where a specific protein binds. They do this with a speed and precision that goes beyond what has been achieved with previous technologies. DNA-binding proteins control transcription, replication, DNA repair, and chromosome segregation. Given the importance of these proteins, identifying their binding sites www.sciencemag.org SCIENCE VOL 316 Published by AAAS throughout the genome has occupied much attention in recent years. The most common method of locating these sites within a living cell is known as chromatin immunoprecipitation (ChIP). In this approach, cells are treated with a reagent, typically formaldehyde, that crosslinks protein and DNA, and then the cells are lysed. Chromatin (the complex of proteins and DNA in chromosomes) is isolated, the DNA is sheared into small fragments, and an antibody is added to precipitate the protein and its associated DNA. The DNA that is liberated after reversal of the protein-DNA crosslinks is then analyzed. In the initial uses of this method, researchers analyzed the DNA to determine whether single genes were enriched by the immunoprecipitation. 8 JUNE 2007 1441
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